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Anatomy and Relationships of Bolong Yixianensis, an Early Cretaceous Iguanodontoid Dinosaur from 19 Western Liaoning, China

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Anatomy and Relationships of Bolong Yixianensis, an Early Cretaceous Iguanodontoid Dinosaur from 19 Western Liaoning, China IU Press uncorrected proof. Copyrighted material. For proofing purposes only. Anatomy and Relationships of Bolong yixianensis, an Early Cretaceous Iguanodontoid Dinosaur from 19 Western Liaoning, China Wu Wenhao* and Pascal Godefroit The skeleton (YHZ-001) of a new basal iguanodontoid was discovered in 19.1. A, Bolong yixianensis, YHZ-001 (holotype). Skull and partially articulated postcranial skeleton, the middle part of the Yixian Formation in western Liaoning, China. as discovered, in right lateral view. Abbreviations: Bolong yixianensis Wu, Godefroit, and Hu, 2010, is characterized by cranial, L, left; R, right. B, Skull reconstruction in left dental, and postcranial autapomorphies, as well as a unique combina- lateral view. tion of characters. A phylogenetic analysis reveals that Bolong is the most primitive Hadrosauroidea described so far. During the Lower Cretaceous, Iguanodontoidea were subdivided into Iguanodontidae, which mainly oc- cupied Neopangean territories, and Hadrosauroidea in Asia. The presence of Iguanodontoidea in the middle part of the Yixian Formation indicates that connections between Asia and western North America and/or Europe were already established during or before the Barremian. Iguanodontoidea (=Hadrosauriformes sensu Sereno, 1997) is defined as Introduction Iguanodon, Parasaurolophus, their most recent common ancestor, and all descendants (Sereno, 1998, amended). During the Early Cretaceous, iguanodontoids had achieved a pan-Laurasian distribution and were also represented in Africa (Norman, 2004). During the Upper Cretaceous, ad- vanced Iguanodontoidea, or Hadrosauridae (a node-based taxon defined as the most recent common ancestor of Bactrosaurus and Parasaurolphus, plus all the descendants of this common ancestor; see Norman, 2004, fig. 11.22), replaced basal iguanodontoids and became the most diverse and abundant large vertebrates of Laurasia during the Campanian and the Maastrichtian. Many new basal iguanodontoids were described during the 2000s, most of which have been recovered from Early Cretaceous deposits in China and Mongolia, including Altirhinus kurzanovi, from Khuren Dukh (late Aptian–Early Albian) of Mongolia (Norman, 1998), “Probactrosaurus” ma- zongshanensis and Equijubus normani, from the Ximinbao Group (Aptian– Albian) of Gansu province (Lü, 1997; You et al., 2003b), Jintasaurus menis- cus, from the Xinminpu Group (?Albian) of Gansu province (You and Li, 2009), Nanyangosaurus zhugeii, from the Sangping Formation (?Albian) of Henan province (Xu et al., 2000), and Penelopognathus weishampeli, from the Bayan Gobi Formation (Albian, Lower Cretaceous) of Inner Mongo- lia (Godefroit et al., 2005). Lanzhousaurus magnidens, represented by an incomplete skeleton from the Hekou Group (Early Cretaceous) of Gansu province, may represent a more basal iguanodontian (You et al., 2005). 293 19-BD Ch19 (292-333).indd 293 2/14/12 8:54 AM IU Press uncorrected proof. Copyrighted material. For proofing purposes only. The Early Cretaceous Jehol Biota of western Liaoning province in China is famous for its abundant, extraordinarily diversified, and exquisitely preserved fossils. The dinosaur fauna of the Jehol Biota is dominated by small-bodied taxa (<3 m in body length), including a variety of coelurosau- rian theropods, basal ceratopsians, the basal ornithopods Jeholosaurus, and the ankylosaur Liaoningosaurus (Xu and Norell, 2006; Zhou, 2006). Larger herbivorous dinosaurs are rare in the Jehol fauna and include the titanosau- riform Dongbutitan dongi (Wang et al., 2007) and the basal iguanodontoid Jinzhousaurus yangi (Wang and Xu, 2001; Barrett et al., 2009; Wang et al., 2011). Shuangmiaosaurus gilmorei, on the basis of an incomplete and de- formed skull, was also collected in western Liaoning province, but in the younger Sunjiawan Formation (late Early or early Late Cretaceous; You et al., 2003a). Here we describe the skeleton of a new basal iguanodontoid collected in 2000 in the middle part of the Yixian Formation to supplement the ini- tial report by Wu et al. (2010). The fossil site is located at Bataigou, Toutai county. This incomplete skeleton was fossilized lying on its left flank, with the limbs roughly perpendicular to the vertebral column and parallel to each other, and with the skull and neck retracted over the back (Fig. 10.1). The estimated length for the complete skeleton (the tip of the tail is miss- ing) is ~4 m (smaller than the holotype of Jinzhousaurus, which is 5–5.5 m). For the sake of convenience, all nonhadrosaurid Iguanodontoidea will be termed “basal iguanodontoids” hereafter. Comparisons are made to other Iguanodontia on the basis of published descriptions of Tenontosaurus spp. (Ostrom, 1970; Forster, 1990; Winkler et al., 1997), Dryosaurus spp. (Janensch, 1955; Galton, 1983), Zalmoxes spp. (Weishampel et al., 2003; Godefroit et al., 2009), Camptosaurus dis- par (Gilmore, 1909; Erickson, 1988), Iguanodon bernissartensis (Norman, 1980), Mantellisaurus atherfieldensis(Norman, 1986), Ouranosaurus nige- riensis (Taquet, 1976), Lanzhousaurus magnidens (You et al., 2005), Lur- dusaurus arenatus (Taquet and Russell, 1999), Nanyangosaurus zhugeii (Xu et al. 2000), Jinzhousaurus yangi (Barrett et al., 2009), Equijubus normani (You et al., 2003b), Altirhinus kurzanovi (Norman, 1998), Penelopognathus weishampeli (Godefroit et al., 2005), Fukuisaurus tetoriensis (Kobayashi and Azuma, 2003), Probactrosaurus gobiensis (Norman, 2002), “Probactosaurus” mazongshanensis (Lü, 1997), Eolambia caroljonesa (Kirkland, 1998), Proto- hadros byrdi (Head, 1998), Bactrosaurus johnsoni (Gilmore, 1933; Godefroit et al., 1998), Levnesovia transoxiana (Sues and Averianov, 2009), Shuangmi- aosaurus gilmorei (You et al., 2003a), Tethyshadros insularis (Dalla Vecchia, 2009), Telmatosaurus transsylvanicus (Weishampel et al., 1993), and various Euhadrosauria. Jintasaurus meniscus You and Li, 2009, was published after the completion of this chapter and has not been included in the analysis. Systematic Paleontology Dinosauria Owen, 1842 Ornithischia Seeley, 1887 Ornithopoda Marsh, 1881 Iguanodontia Dollo, 1888 [Sereno, 1986] Iguanodontoidea Cope, 1869 Hadrosauroidea Cope, 1869 294 Wu and Godefroit 19-BD Ch19 (292-333).indd 294 2/14/12 8:54 AM IU Press uncorrected proof. Copyrighted material. For proofing purposes only. 19.2. Bolong yixianensis, YHZ-001 (holotype). Rostral portion of the skull in left lateral view. Anatomy and Relationships of Bolong yixianensis 295 19-BD Ch19 (292-333).indd 295 2/14/12 8:54 AM IU Press uncorrected proof. Copyrighted material. For proofing purposes only. Bolong yixianensis Wu, Godefroit, and Hu, 2010 (Figs. 19.1–19.11) Etymology. The generic name is in honor of Bo Haichen and Bo Xue, who discovered and excavated the holotype; long means “dragon” in Chi- nese. The specific name refers both to the Yixian Formation, where the holotype was discovered, and to the city of Yixian, where the holotype is housed and displayed. Holotype. YHZ-001, housed in Yizhou Fossil Museum, Yixian City, Liaoning province (P.R. China). Locality and horizon. Bataigou, Toutai, Yixian County, western Lia- oning province, P.R. China. GPS coordinates: N41°36'6.79", E121°7'43.1". Dakangpu Member (equivalent to the Dawangzhangzi Beds) of the middle part of Yixian Formation, Late Barremian–Early Aptian (Smith et al., 1995; Swisher et al., 1999, 2002). Diagnosis. Differs from all other iguanodontoid taxa in possessing the following autapomorphies: depressed area at the junction between maxilla and lacrimal, corresponding to the position of the antorbital fenestra in Iguanodon, Mantellisaurus, and Ouranosaurus; caudal ramus of prefrontal forming a rostrocaudally depressed area above the orbital margin; ventral process of predentary extending caudally parallel to the ventral margin of the predentary main body; rostrodorsal articular surface for the predentary occupying less than two-thirds of the height of the rostral part of the dentary and rostral tip of dentary therefore situated above the ventral third of the bone; primary ridge deflected distally on maxillary crowns; ulna and radius proportionally short and robust (ratio craniocaudal height of proximal part/ length = 0.28 for ulna and 0.33 for radius); proximal and distal ends of ra- dius nearly symmetrically enlarged craniocaudally and triangular in medial and lateral views; postacetabular process of ilium dorsoventrally narrow (ratio length/maximal height = 2); metatarsals proportionally short: ratio metatarsal III/femur = 0.18. Description: All the measurements taken on YHZ-001 are compiled in Appendix 19.1. The preorbital region of the skull is well preserved, and both sides were prepared. Like the rest of the skeleton, it was laterally crushed dur- ing fossilization. Bones are usually preserved in natural connection on the left side of the skull (Fig. 19.2), in direct contact with the sediment after the death of the animal, whereas they were slightly displaced on the right side of the skull (Fig. 19.3). A fault bisects the skull, and the fronto-orbital region of the skull has consequently been lost. Only the right side of the back of the skull has been prepared (Fig. 19.4), but this region is crushed and poorly preserved. External naris. The external naris is drop shaped and proportionally larger (45% of preorbital length) than in Jinzhousaurus (38%), Iguanodon, Mantellisaurus (approximately 28–30%), and Ouranosaurus (approximately 18%). However, it is proportionally much shorter than in Altirhinus (ap- proximately 56%) and Hadrosauridae. The
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